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The Porifera represent one of the only two recent nerveless and muscleless metazoan phyla. Nevertheless, sponges provide behavioral, physiological, pharmacological, morphological, and, more recently, an increasing amount of genetic evidence for a paracrine pre-nervous integration system. Although this system might be derived, it allows us to draw conclusions, on the basis of comparative data, about the origin of the nervous system sensu stricto as found in the eumetazoan phyla. The goal of the present review is to compile recent evidence on the sponge integration systems. Based on this framework, new light is also shed on the evolutionary origin of the eumetazoan synaptic nervous systems, which can be regarded to form an evolutionary biochemical continuum with the paracrine signaling system in sponges. Thus, we can assume that the evolutionary transition from a paracrine-dominated, pre-nervous system to an electrochemically dominated, primordial nervous system resulted in part from compartmentalization effects. As intermediate evolutionary stages, regionalized synapse precursor areas might have occurred within pre-nervous cells, which foreshadowed the highly organized synaptic scaffolds present in recent nerve cells of the Eumetazoa.
Cycliophora is one of the most recently described metazoan phyla and hitherto includes only two species: Symbion pandora and Symbion americanus. With a very complex life cycle, cycliophorans are regarded as an enigmatic group with an uncertain phylogenetic position, although they are commonly considered lophotrochozoan protostomes. In order to extend the database concerning the distribution of immunoreactive substances in the free-swimming chordoid larva of S. pandora, we investigated synapsin immunoreactivity using fluorescence-coupled antibodies in combination with confocal laserscanning microscopy. Moreover, we analyzed the co-localization patterns of synapsin, serotonin, and RFamide-like immunoreactivity in the chordoid larva by 3D imaging technology based on the confocal microscopy image stacks. Synapsin is expressed in large parts of the bilobed anterior cerebral ganglion including anterior and dorsal projections. Two pairs of ventral neurites run longitudinally into the larval body of which the inner pair shows only weak, scattered synapsin immunoreactivity. In addition, a lateral synapsin immunoreactive projection emerges posteriorly from each ventral longitudinal axon. Double immunostaining shows co-localization of synapsin and serotonin in the cerebral ganglion, the outer and the inner ventral neurites, and the anterior projections. Synapsin and RFamide-like immunoreactivity co-occur in the cerebral ganglion, the outer ventral neurites, and the dorsal projections. Accordingly, the cerebral ganglion and the outer ventral neurites are the only neural structures that co-express the two neurotransmitters and synapsin. The overall neuroanatomical condition of the cycliophoran chordoid larva resembles much more the situation of adult rather than larval life cycle stages of a number of spiralian taxa.
The topic of tissue and organ regeneration has been of interest to life scientists ever since the phenomenon was noticed. The reason for this is obvious: if one can learn what drives and controls regeneration, i.e., how lost or damaged structures can be replaced, one not only has a better chance to understand an animal's embryogenesis and evolutionary relationship with other taxa, but one would also be in a better position to treat organ loss or tissue damage in humans. In this context, the possible restitution of individual sensory neurons or nerve projections has been of special interest to us. We identified central visual projections in several gastropod species and found that: (1) projections are very extensive across the brain and (2) they have connections with other systems and organs (including, most likely, non-ocular skin photoreceptors) that may be involved in the integration of signals from different sensors. Investigations of afferent and efferent visual elements at a morphological level should help reveal the neuronal basis of a gastropod's behavioral reactions.
The nervous system of the meiobenthic priapulid species Tubiluchus troglodytes is described by immunohistochemistry and confocal laser scanning microscopy. The brain is circumpharyngeal, consisting of a central ring of neuropil and both anterior and posterior somata. From the brain emerges a ventral nerve cord, which shows ganglion-like swellings in the neck and caudal region. The introvert includes longitudinal neurite bundles running below and between the rows of scalids, with a small cluster of sensory cells under each scalid. In the body wall of the neck and trunk region, longitudinal and circular neurite bundles are present in an orthogonal pattern. The tail is innervated from the caudal swelling of the ventral nerve cord; it also includes longitudinal and circular bundles in an orthogonal pattern. The pharynx has a reticulated system of neurite bundles running between the pharyngeal teeth and fimbrillae. Below each tooth and fimbrilus is a ganglion-like cluster of somata. The intestine is surrounded by a nerve net. The data on the nervous system are compared within other priapulids and with other species of Scalidophora (Kinorhyncha and Loricifera).
The formation of the central nervous system of the stomatopod crustacean Gonodactylaceus falcatus is described by means of antibody stainings against synapsin and α-tubulin. It is shown that the longitudinal fiber tracts of the ventral nervous system are formed by two centers of origin comprising a number of pioneer neurons, one at the posterior part of the forming brain, the other in the area of the telson anlage at the posteriormost region of the embryo. In addition to the lateral anlagen of the connectives, a median longitudinal nerve is formed beginning in the mandibular segment neuromere. In contrast to those of other segments, the mandibular ganglia are connected by a single commissure. The brain forms a circumoral ring. There is evidence that the deutocerebrum possesses praestomodeal and poststomodeal commissural fibers. The anlage of the nauplius eye reveals a specific pattern of pigment and sensory cells with the two pigment cells expressing synapsin. Clear differences between the expression patterns of synapsin and α-tubulin recommend the combination of a variety of antibodies to gain a complete picture of embryonic neuroanatomy. Our results show overall similarities to other malacostracan and non-malacostracan crustaceans. The comparisons with other crustaceans and arthropods indicate homology of crustacean nauplius eyes, a circumoral deutocerebrum, and a more widespread occurrence of posterior pioneer neurons forming the axon scaffold of the ventral central nervous system than previously thought.
We examined the brain architecture in different species of Chaetognatha using immunofluorescence methods with a set of nervous system markers and confocal laser-scan microscopic analysis. These markers include antibodies against synaptic proteins, RFamide-related peptides, and tyrosinated tubulin, as well as a marker of cell nuclei. Furthermore, we present a 3D reconstruction based on histological section series. Our results expand the previous knowledge on neuroanatomy in Chaetognatha. We suggest a structural and functional subdivision of the rather complex chaetognath brain into two domains, a posterior domain that may be primarily involved in the integration of sensory input, and an anterior domain that may be involved in the control of the mouthparts and the anterior part of the digestive system. Immunolocalization of a neuropeptide suggests the presence of an identifiable group of neurons associated with the brain of all species examined here. However, our data also reveal a certain degree of interspecific variation and divergence within the Chaetognatha concerning, for example, the pattern of nerves branching off the brain and the proportional sizes of the various neuropil compartments. We compare our data to brain architecture in various other representatives of Protostomia and Deuterostomia. The chaetognath brain fits within the range of structural variation encountered in protostomian brains, and we cannot find any brain characteristics that would argue in favor of placing chaetognaths outside of the Protostomia. Rather, we see the circumoral arrangement of their cephalic nervous system as an argument that suggests protostome affinities.
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